Longhorn beetles (Coleoptera, Cerambycidae)

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Christian Cocquempot, Ake Lindelöw

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Christian Cocquempot, Ake Lindelöw. Longhorn beetles (Coleoptera, Cerambycidae). Alien terrestrial arthropods of Europe, 4 (1), Pensoft Publishers, 2010, BioRisk, 978-954-642-554-6. �10.3897/biorisk.4.56�. �hal-02823535�

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Longhorn beetles (Coleoptera, Cerambycidae). Chapter 8.1 193

Longhorn beetles

(Coleoptera, Cerambycidae)

Chapter 8.1

Christian Cocquempot1_{, Åke Lindelöw}2

1 INRA UMR Centre de Biologie et de Gestion des Populations, CBGP, (INRA/IRD/CIRAD/Montpellier

SupAgro), Campus international de Baillarguet, CS 30016, 34988 Montférrier-sur-Lez, France 2 Swedish university of agricultural sciences, Department of ecology. P.O. Box 7044, S-750 07 Uppsala, Sweden

Corresponding authors: Christian Cocquempot (cocquemp@supagro.inra.fr), Åke Lindelöw (

Ake.Linde-low@ekol.slu.se)

Academic editor: David Roy | Received 28 December 2009 | Accepted 21 May 2010 | Published 6 July 2010

Citation: Cocquempot C, Lindelöw Å (2010) Longhorn beetles (Coleoptera, Cerambycidae). Chapter 8.1. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(1): 193–218. doi: 10.3897/biorisk.4.56

Abstract

A total of 19 alien longhorn beetle species have established in Europe where they presently account for ca. 2.8 % of the total cerambycid fauna. Most species belong to the subfamilies Cerambycinae and Laminae which are prevalent in the native fauna as well. Th e alien species mainly established during the period 1975–1999, arriving predominantly from Asia. France, Spain and Italy are by far the most invaded countries. All species have been introduced accidentally. Wood-derived products such as wood- packaging material and palettes, plants for planting, and bonsais constitute invasive pathways of increasing impor-tance. However, only few species have yet colonized natural habitats outside parks and gardens. Present ecological and economical impacts, and future trends are discussed.

Keywords

Cerambycidae, Europe, Introductions, Establishments, Biogeographical origins, Pathways, Impacts

8.1.1 Introduction

Th e coleopteran family Cerambycidae (longhorn beetles) is currently classifi ed in the superfamily Chrysomeloidea, along with the families Vesperidae and Distenidae (Hunt et al. 2007, Szeoke and Hegyi 2002). Cerambycidae is a large family comprising about BioRisk 4(1): 193–218 (2010)

doi: 10.3897/biorisk.4.56 www.pensoftonline.net/biorisk

Copyright C. Cocquempot, Å. Lindelöw. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

RESEARCH ARTICLE

BioRisk

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40000 described species worldwide. Longhorn beetles are all phytophagous. Larvae may be found in conifer, deciduous and fruit trees, in bushes and herbaceous plants. Th ey are mainly xylophagous borers of living, decaying or dead wood. Some species also bore small twigs, roots or fruit endocarps. Th ey usually have a long period of lar-val development, some species being capable of developing in woody material a long time after the death of the tree. Th ey are thus very susceptible to transport with wood products, facilitating their introduction and establishment.

Th e oldest known introduction of a longhorn beetle from one continent to anoth-er was probably that of the house boranoth-er, Hylotrupes bajulus (L., 1758), which was fi rst described by Linnaeus from both Europe and ‘America septentrionali’ (von Linnaeus 1758). Since a study by Duff y in 1953 (Duff y 1953a) for Great Britain, there has been no further large synthesis of the alien cerambycid species introduced to Europe. Since 1999, the development of research interests in the Asian longhorn beetles, Anoplophora spp., in North America has raised awareness of the risks presented by cerambycid im-portation and provided a baseline for subsequent studies (Haack et al. 2000, Haack et al. 2010). Th ere is an urgent need for a comprehensive literature review of the alien cerambycids that have successfully established in Europe.

Th e exponential growth in the volume of international trade in both horticulture and forestry has allowed an increasing number of wood products and ornamental plants potentially containing cerambycids to arrive in Europe. More than 250 species have been introduced to Europe or moved within Europe since the middle of the 18th cen-tury (Cocquempot 2007) but most of them never established. We have identifi ed 19 species alien to Europe that have established in Europe but have not yet been eradicated.

8.1.2 Taxonomy of the Cerambycid species alien to Europe

Taxonomy in Cerambycidae sensu lato is not well established (e.g., Hunt et al. 2007, Lawrence and Newton 1995, Napp 1994, Özdikmen 2008, Sýkorová 2008) but a gen-eral consensus exists about the presence in Europe of 7 subfamilies, namely Ceramby-cinae, Lamiinae, Lepturinae, Necydalinae, Prioninae, Spondylidinae, and Vesperinae (the latter being sometimes considered as a valid family). A total of 677 native species are known to occur in Europe (Althoff and Danilevsky 1997, Fauna Europaea), being largely dominated by 3 subfamilies (Lamiinae- 343 spp.; Cerambycinae- 158 spp.; Lepturinae- 130 spp.) which account for 93.2% of the total.

Th e 19 alien species established in Europe belong to only 3 of these subfamilies, Cerambycinae, Laminae and Prioninae (Table 8.1.1). Th e alien species are mostly rep-resented by the subfamily Cerambycinae, followed by Lamiinae but the relative propor-tion of aliens compared to the total cerambycid fauna is still limited (<6%) in these two subfamilies. By contrast, the proportion of aliens is much more important in Prioninae with 2 species adding to 10 native ones (Fig. 8.1.1.). In addition, Parandrinae, a sub-family which is not represented in the native European entomofauna, is represented by Parandra brunnea, a North American species introduced in Germany (Nüssler 1961).

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Two more alien species have been introduced and established in Israel, Batocera

rufomaculata (DeGeer, 1775) (Bytinski-Salz 1956, Chikatunov et al. 1999, Sama et

al. 2010) and Xystrocera globosa (Olivier, 1795) (Chikatunov et al. 2006, Sama et al. 2010), but they have not yet spread to Europe and were not considered in Table 8.1.1.

Table 8.1.2 gives a list of species of European origin introduced through human activity in another part of Europe (aliens in Europe). Th ese species are mostly of Medi-terranean origin introduced in more northern areas and species from Continental Eu-rope introduced to the Atlantic islands.

8.1.3 Major biological characteristics of the cerambycid species alien to Europe

Lepturinae but also Prioninae and Parandrinae share some biological characteristics that reduce their probability of introduction. Larvae in these subfamilies develop in decaying wood and are rarely imported with wood products or living plants. Intercep-tions have shown that they are mainly introduced through accidental importation in industrial packages or in stocks of perishable vegetables. Only a few species of Lepturi-nae (Tribe Rhagiini, and some Lepturinii) developing on recently felled trees are likely to be successfully introduced through the wood trade. Th e importation of living potted plants is also a potential new pathway for Prioninae.

Cerambycinae and Lamiinae seem more predisposed to introduction. Most species develope in living plants and several Cerambycinae undertake their entire life-cycle in dead wood, e.g. the cosmopolitan tribe Hesperophanini and the species Hylotrupes

bajulus and Gracilia minuta. Th us, Cerambycinae and Lamiinae can easily survive Figure 8.1.1. Relative importance of the subfamilies of Cerambycidae in the alien and native entomo-fauna in Europe. Subfamilies are presented in a decreasing order based on the number of alien species. Species alien to Europe include cryptogenic species. Th e number over each bar indicates the number of species observed per family.

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throughout the importation process of living plants including bonsai (e.g.

Anoplo-phora chinensis (Cocquempot 2007, EPPO 2006, van Rossem et al. 1981, Schmidt

and Schmidt 1990)), recently felled logs and other non-aged wood products (e.g.

Ano-plophora glabripennis (Cocquempot et al. 2003, Haack et al. 2000), Monochamus spp.

(Cocquempot 2007, Cocquempot (Unpubl.), Duff y 1953a), Chlorophorus annularis (Cocquempot 2007) and Phoracantha spp. (Cocquempot and Debreuil 2006)). Spe-cies in the genera Hesperophanes, Trichoferus, and Stromatium can emerge from wood products even several years after importation (Duff y 1953a).

Once a population is introduced, the capability for natural dispersal constitutes an important factor for establishment success. Although our knowledge about the disper-sal behaviour of alien longhorn beetles is still rather limited and mostly concerns only a few species of recent invaders such as Anoplophora glabripennis (Smith et aol. 2001) and A. chinensis (Adachi 1990, Komazaki and Sakagami 1989), this variable is impor-tant when designing an eradication attempt (MacLeod et al. 2002).

8.1.4 Temporal trends of introduction in Europe of alien Cerambycids

Figure 8.1.2 presents the temporal changes in the records of Cerambycid species alien

to Europe from 1492 to 2007. Cerambycids have tracked trade routes since the

begin-ning of overseas communications. Th e fi rst species to have moved are those which live in dry wood and undergo a long stage of larval development. Th ese species have be-come cosmopolitan (e.g. Hylotrupes bajulus) or nearly so (e.g. Stromatium spp.). With the increased speed of international transport from 1850 to 1925, species with shorter life cycles were able to reach Europe alive and become established, e.g. Neoclytus

acu-minatus (Reineck 1919, Sama 2002, Tassi 1969). Later, only two species were

intro-duced from North America to Europe via the US eff ort to supply extra furniture and increase military material after the 1st_{World War (i.e., Parandra brunnea, Neoclytus}

acuminatus). Subsequently, 50 years passed until a second wave of introduction

ar-rived alongside with the rapid development of international exchange of goods and transportation after the 2nd_{World War. During the recent period, two further species} have been detected in the wild - Anoplophora chinensis in 2000 in Italy (Colombo and Limonta 2001) and A. glabripennis in 2001 in Austria (Dauber and Mitter 2001).

Th e number of interceptions of Cerambycids is still increasing throughout Eu-rope. However, more eff ective control at borders is like to have reduced establishments following interception or introductions. Th e importation of exotic plants also off ers opportunities for introduction but also constraints the establishment of some alien species. For example, Phoracantha spp. could not have been introduced without the importation and mass cultivation of its host plants, Eucalyptus spp. in the Mediter-ranean basin. In south-eastern France, an Australian cerambycid, Bardistus cibarius (Newman, 1841) could survive only on its original host plant, an introduced grass tree (Xanthorrhoea sp., Xanthorrhoeaceae); the beetle population disappeared immediately after the infested host plants were removed (Cocquempot 2007). Th e case of Batocera

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rufomaculata (DeGeer, 1775) found in Munster’s Zoo (Germany) is similar

(Coc-quempot 2007) although this tropical species has established in Israel since at least 1948 (Bahillo de la Puebla and Iturrondobeitia-Bilbao 1995, Plavilstshtikov 1934, Sama et al. 2010). Th e combination of importation of longhorn beetle species with their specifi c host plant or groups of plants followed by establishment is rare. However the establishment of A. chinensis is an exception. Other species are frequent intercepted at border controls, e.g. Mimectatina meridiana (Matsushita, 1933) with Cycas fruits from Japan (Cocquempot 2007) or Trichoferus campestris (Faldermann 1835) with

Sa-lix timber from China (Cocquempot 2007).

Th e degree of polyphagy is also an important factor in the likelihood of establish-ment. Polyphagous species appear to have a higher potential to establish than oligopha-gous and monophaoligopha-gous species. Th e large number of hosts utilised by Anoplophora spp. (Cocquempot et al. 2003, Hérard and Roques 2009, Maspero et al. 2007a) is a main factor in the diffi culty in eradicating this species for example. Th ese diffi culties appear much less important for oligophagous species such as Callidiellum rufi penne (Bahillo and Iturrondobeitia-Bilbao 1995, Campadelli and Sama 1988, Plavilstshtikov 1934) Figure 8.1.2. Temporal changes in the mean number of new records per year of Cerambycid species alien to Europe from 1492 to 2007. Th e number over each bar indicates the absolute number of species newly recorded per time period.

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or Phoracanthine species. It is also the case for the North American wood borer Saperda

candida (Fabricius, 1787), which was introduced in Germany in 2008 but apparently

did not established yet (EPPO 2008, Nolte Krieger 2008). By contrast, Monochamus species have a regime close to polyphagy, including a large number of conifer species, and may spread throughout Europe. Th ere is no example of establishment in Europe of a strictly monophagous exotic long-horned beetle. Species with a limited host range do not seem to be capable of going beyond the interception or introduction stage, e.g.

Bardistus cibarius (Cocquempot 2007).

8.1.5 Biogeographic patterns of the cerambycid species alien to Europe

Alien species established in Europe mostly originated from Asia, followed by Africa (Figure 8.1.3). Th e region of origin appears to depend on the major trade routes de-veloped by each country. Some North African species have colonized Mediterranean countries such as Spain, France, and Malta for example. Other African species have often been intercepted but only Phryneta leprosa has established in Malta where the climate is favourable for development (Mifsud and Dandria 2002). Long-established trade routes between Iberian countries and South American countries have resulted in some historic, isolated establishments in the Spanish and Portuguese Atlantic Is-lands but with a limited risk of further expansion (Lemos-Perreira 1978, Méquignon 1935). With the numerous interceptions in the U.K (Duff y 1953a) together with the colonial trade routes with African and Asiatic countries, it is surprising that only

Tri-nophylum cribratum has established to date (Gilmour 1948); the incompatible climate

may negate the development of tropical and subtropical species. Two species native to North America, Parandra brunnea and Neoclytus acuminatus, also colonized Europe at the beginning of the last century. Th e fi rst species is well established but restricted to Dresden (Germany) (Nüssler 1961). Th e second is widely established in the Mediter-ranean area but its populations appear to be declining (Brustel et al. 2002). Beside these two species, there have been no further establishments originating from North America; the pathway of transported material is mainly in the reverse direction, from Europe to America.

Some Australian species have reached Europe but only those using Eucalyptus

(Pho-racantha spp.) have successfully established (Cocquempot and Sama 2004) and only in

areas newly planted with these fast-growing tree species. Th e large diff erences in spe-cies composition between the fl oras of Australia and Europe probably accounts for the failure of Australasian longhorn beetles such as in Bardistus cibarius on Xanthorrhoea sp. (Cocquempot 2007) to establish.

Recent increases in commercial traffi c from Asia (especially China) to Europe has accounted for the introduction of a number of new species of cerambycids. Striking examples are Callidiellum rufi penne which has recently established in Spain (Bahillo de la Puebla and Iturrondobeitia-Bilbao 1995) and Italy (Campadelli and Sama 1988),

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not eradicated in several countries (Hérard and Roques 2009, Maspero et al. 2007a),

Psacothea hilaris (Pascoe, 1857) under eradication in Italy (Cocquempot 2007, Jucker

et al. 2006), and Monochamus alternatus Hope, 1842 intercepted a number of times in Germany (Cocquempot 2007) and France (Cocquempot Unpubl.) but not yet estab-lished. A fi nal case, Xylotrechus stebbingi, is less clear. It is believed that an initial intro-duction from its native area of central Asia to Asia Minor was followed by a step-wise expansion into southern Europe and North Africa (Cocquempot and Debreuil 2006, Sama 2002, Šefrová and Laštůvka 2005).

Alien cerambycid species are not evenly distributed throughout Europe. Large dif-ferences in the number of aliens are apparent between countries, France, Italy and Spain being by far the most invaded (Figure 8.1.4).

8.1.6 Main pathways of introduction to Europe of alien cerambycid spe-cies

All alien longhorn beetles established in Europe have been introduced accidentally; there are no examples of a successful, deliberate introduction. Th e principal pathways of arrival have been identifi ed and presented by Frank 2002 and each relates to the import of immature stages that subsequently emerge as adults. Th ere are relatively few records of living adults imported with vegetables or fruits although Eucalyptus beetles,

Phoracantha recurva, were found in a cluster of bananas (Bosmans 2006).

Th e longest established pathway is timber importation for house construction

(Hy-lotrupes bajulus) or building furniture (e.g. Trichoferus spp., Stromatium spp. and Chlo-rophorus annularis arriving with bamboo- made objects (Cocquempot 2007)). Species

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introduced through this pathway have traditionally required a long life cycle but more rapid travel now enables the introduction of species with a one year life cycle. Th e second pathway is via the importation of timber for pulp (e.g., for Phoracantha spp.). A third, more recent, pathway concerns wood packages, palettes and other wood-de-rived products (e.g., for Anoplophora glabripennis) (Hérard and Roques 2009). Th e fi nal pathway is the importation of plants for planting in nurseries, including the bon-sai industry, which has resulted in the arrival of species such as Anopolophra chinensis (Cocquempot 2007, EPPO 2006, van Rossem et al. 1981, Schembri and Sama 1986),

Callidiellum rufi penne and Bardistus cibarius.

All pathways are still prevalent but they vary in importance. Most recent inter-ceptions (from the end of the 20th_{Century) have related to wood-manufactured} products (e.g. Chlorophorus annularis and Trichoferus campestris). Importation of

Eu-calyptus wood for pulp has also resulted in the introduction of a second species of Phoracantha, P. recurva (Miquel 2008). If such importations continues a number of

Figure 8.1.4. Comparative colonization of continental European countries and islands by Cerambycidae species alien to Europe. Archipelago: 1 Azores 2 Madeira 3 Canary islands.

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additional species of this genus, which are mainly related to Eucalyptus(Wang 1995), are expected to arrive.

Since their fi rst usage, wood packaging and palettes have constituted an impor-tant introduction pathway. Th e source material spends suffi cient time as logs without sanitary controls to be colonized by longhorn beetles. When the wood is turned into packages or palettes, infestation occurs mainly as unnoticed early stages (eggs or fi rst- instar larva). Development continues in the woody material during importation and emergence of adults occurs often unnoticed in warehouses, weeks or months after ar-rival. Th is is the case for A. glabripennis, P. hilaris and M. alternatus which may already complete their entire lifecycle before the source wood is processed or destroyed. Wood package is often produced using low quality timber often colonized by longhorn beetle species, which is increasing its potential as a vector.

Other, less signifi cant, introduction pathways have also been identifi ed, yet they typically only transported one or a few individuals which fail to establish. Th e intro-duction route is unknown for other species such as Acanthoderes jaspideus (Méqui-gnon 1935), Oxymerus aculeatus (Alluaud 1935), Deroplia albida, and Phryneta leprosa (Mifsud and Dandria 2002) but they may be related to the uncontrolled importation of wild plants. Natural range expansion cannot be ruled out for a few species which have a nearby native range, e.g. Lucasianus levaillantii (Mayet 1905, Pellegrin and Cocquempot 2001) and Xylotrechus stebbingi (Šefrová and Laštůvka 2005) originating from North Africa and the Middle East, respectively.

8.1.7 Ecosystems and habitats invaded in Europe by alien cerambycid species

Although all natural or artifi cial terrestrial ecosystems and anthropogenic areas which contain trees, bushes or wood products are potentially occupied by alien longhorn bee-tles, establishment in Europe is concentrated in man-made habitats to date, especially in parks and gardens (Figure 8.1.5). To date, only the two clytine beetles, Neoclytus

acumi-natus and Xylotrechus stebbingi, have colonized natural habitats. X. stebbingi is very

com-mon on Eucalyptus cut wood in Crete (Sama 2002) for example and may be related to the polyphagous nature of these two species. Other polyphagous species such as Anoplophora spp. also have the potential to live in urban areas, in cultivated lanes (e.g. planted with poplars) as well as in natural forests where potential host plants occur. However, disper-sal from man-made habitats to natural forests appears to be a slow process. For the fi rst twenty-two years since its arrival in North America, A. glabripennis has been restricted to trees in urban areas until 2008 when it was found in natural forests dominated by Acer trees (Haack et al. 2010). Although such a process has not yet been observed in Europe, there is a strong risk that Anoplophora spp. will spread to naturally-forested landscapes, if the ongoing eradication attempts in Austria, Germany, France and Italy are unsuccessful.

Th e expansion of oligophagous species is inevitably more dependant on the pres-ence of suitable host plants. Th ose using largely- planted trees can spread more easily.

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Th us, Phoracantha spp. that live only in eucalypt trees have colonized ornamental tree plantations in urban areas as well as old plantations such as those found on the Medi-terranean islands and in neighbouring countries, and industrial plantations created for paper pulp. Other established species mostly have a distribution restricted to Mediter-ranean and Atlantic islands. In these areas, anthropogenic ecosystems are mainly colo-nized. A species of considerable concern with conifer forests is Monochamus alternatus, which could potentially become established in coniferous plantations and forests and subsequently transfer the pine wood nematode (Bursaphelenchus xylophilus Steiner & Buhrer, 1934).

8.1.8 Ecological and economic impact of alien cerambycid species

Although there is concern about the potential ecological impact of the invasive long-horn beetles N. acuminatus and X. stebbingi, there is no measure of their impact on trees or any estimation of possible competitive displacement of the native fauna. Th e ecological impact of Anoplophora species may also be important if they establish in European forests. Anoplophora could compete with other arthropods occupying the same niche, but they also create niches for other arthropods that live in tunnels in decaying wood or compete with other saproxylic beetles. Th e joint introduction Figure 8.1.5. Main European habitats colonized by the established alien longhorn beetles. Th e number over each bar indicates the absolute number of alien longhorn beetles recorded per habitat. Note that a species may have colonized several habitats.

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and establishment of the Citrus longhorn beetle, A. chinensis, and its parasitoid,

Aprostocetus anoplophorae Delvare, 2004, exemplifi es the potential risk of adaptation

of imported parasitoids which themselves might not specialise on the native fauna (Delvare et al. 2004).

Although the ecological niche occupied by an alien species may be vacant there remains a risk of secondary infection resulting from their damage. For example, sec-ondary infestation by the pine wood nematode vectored by Monochamus spp. (Evans et al. 2008, Kawai Miho et al. 2006) may cause serious impacts to coniferous trees in all landscapes. M. alternatus has only been intercepted in Germany and France (Cocquempot 2007, Cocquempot (Unpubl.)); yet the pine wood nematode which it vectors was recorded from Portugal in 1999 (Mota et al. 1999). After having been contained for several years in a limited area, the nematode has spread throughout Por-tugal, as well as being eradicated following incursions into Spain in 2008 and Madeira in 2009. A novel association with the native species, M. galloprovincialis (Villiers 1967) has also been reported. Th e expansion as well as new introductions of the pine wood nematode could potentially have a substantial level of economic impact in all areas of coniferous cultivation in Europe.

Other economic impacts are mainly associated with ornamental trees in urban areas, cultivated trees such as poplars and eucalypts and nurseries, including these for bonsai production. Studies of Anoplophora glabripennis in North America and A.

chinensis in China indicate the possible scale of economic damage following

estab-lishment of these species in a new country or in a plantation, especially of poplar or Citrus trees (Cocquempot et al. 2003, Haack et al. 2010, MacLeod et al. 2002). As a control measure, ornamental trees colonized by invasive longhorns must be elimi-nated without consideration of their aesthetic value. Eradication measures entail high costs to be borne by local communities or private owners. Special attention is paid to A. chinensis necessitating complete removal of trees, including the rootstock (Haack et al. 2010).

Poplars or eucalypt plantations can be highly aff ected as has already been the case in China (A. glabripennis on poplars) and in Spain (Phoracantha spp.), where infested trees become unsuitable for pulp and wood exploitation. Th e Citrus longhorn beetle is also considered as an important risk for all Citrus fruit production in the Mediter-ranean area and its islands.

Th e nursery industry is already concerned. Th ere are several examples of introduc-tions or establishments of potentially invasive species such as Callidiellum rufi penne and Anoplophora chinensis, with the imports of nursery plants. Nurseries can them-selves be vectors of aliens when they dispatch their products.

Th e eradication process established for quarantine species aims to limit introduc-tions although only a few eradicaintroduc-tions have been offi cially reported in Europe, e.g. as for Anoplophora chinensis in France (Hérard et al. 2006, Hérard and Roques 2009). Phytosanitary interceptions at borders are likely to have prevented a number of intro-ductions and further establishments (e.g., Monochamus alternatus, Trichoferus

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(Cocquem-Figure 8.1.6. Adults of some alien longhorn beetle species. a Phoracantha semipunctata b Phoracantha recurva c Mimectatina meridiana (Credit: Christian Cocquempot) d Xylotrechus stebbingi (Credit: Vítěslav Maňák) e Bardistes cibarius (Credit: Christian Cocquempot) f Psacothea hilaris g Parandra brunnea (a, b, e, f, g: Credit: Henri-Pierre Aberlenc).

pot 2007) whilst at the same time, several non-quarantine species not submitted to importation controls have become established (e.g., Xylotrechus stebbingi, Phoracantha

semipunctata, Neoclytus acuminatus). Th is illustrates the importance of quarantine spe-cies lists, which should be preventive and not only curative to be most eff ective.

Human-mediated dispersal should also be tightly controlled during the eradica-tion process. Without due respect for control obligaeradica-tions, eradicaeradica-tion can fail. For example, the long delay by Italian authorities in applying control measures and strong management measures against Anoplophora chinensis (EPPO 2009, Jucker et al. 2007) or inadvertent movement of untreated wood material for A. glabripennis in New-York (Haack et al. 1997) are examples of ineff ective eradication effi cacy.

a b f

e g

c

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8.1.9 Expected trends

Th e combination of increasing volumes of trade, the increased speed of import of potential vectors, the diversity of sources and sites for introduction is likely to result in increasing invasion risk (Cocquempot 2007). All recently established species alien

to Europe have been intercepted too late after their introduction and have been

out-side offi cial institutional controls. Th ese factors make it increasingly diffi cult for rapid eradication after initial arrival. Eff ective monitoring of each point of possible entry is unfeasible when the key pathways identifi ed here have diff erent vectors and locations of arrival (e.g. airports, harbours, stations, lorry parks), and there are major diff er-ence in the quality of phytosanitary controls between European countries, particularly following the enlargement of the EU. Th e risk depends on volume and diversity of vector material imported, and subsequently there is greatest risk in countries such as the UK, France, Spain, Italy, Netherlands, Belgium and Germany. Th e case of

Anopolo-phora glabripennis in North America and Europe clearly demonstrates the possibility of

spread in our continent; such detailed assessment is required for all potentially invasive longhorn beetles (MacLeod et al. 2002).

According to Worner (2002), progress in the knowledge of invasion processes and associated preventive measures have not been followed by actions since the late 1980’s. Preventive methods are still routinely applied, e.g. the application of ISPM 15 (Interna-tional Standard for Phytosanitary Measures No.15), which set standards for heat treat-ment and fumigation of wood product materials used in international trade is likely to limit the arrival of longhorn beetles related to these materials although a few have been found to survive (Haack et al. 2010). However, this method is not uniformly applied to all imported living trees, shrubs plants for planting or bonsais. Th us, a high number of imported bonsais or other nursery trees infested with Anoplophora chinensis are still discovered (Hérard and Roques 2009). Although importation controls could be im-proved, they will never off er full protection. Further, controls which reduce the risk of introduction are mainly restricted to quarantine species. Post-interception or controls at importation points should be extended to all the potential pests posing risk and not be restricted to quarantine species already intercepted, introduced or established.

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Wollaston TV (1863) On the Canarian Longicorns. Journal of Entomology 8: 99–110. Worner SP (2002) Predicting the invasive potential of exotic insects. In: Hallman GJ, Schwalbe

CP Invasive Arthropods in Agriculture: Problems and Solutions. Enfi eld, UK: Sciences Pub-lishers Inc., 119–137.

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F amily _Species Status R egime N ativ e range 1st r ecor d in E u rope In vaded countries H abitat H osts R efer ences A canthoder es jaspidea G ermar , 1824 A phyto -phagous B razil 1880, PT -AZ O P T -AZ O I2 A cacia, _Albizzia

Borges et al. 2005, Méquignon 1935, S

errano 1982 A cr ocinus longimanus (Linnaeus, 1758) A phytophagous B razil 1977, PT PT , PT -MAD I2 M oraceae, A pocynaceae Lemos-P err eira 1978 , V iv es 1995 A noplophor a chinensis (F örster , 1848) (= A . malasiaca Th ompson, 1865) A phytophagous China S outh-Central 2000, IT IT , NL FB, F A, I2, G A cer , B etula, C arpinus C itr us, Cor ylus, R osa and deciduous shr ubs (polyphagous)

Cocquempot 2007, Colombo and Limonta 2001, 2009a, EPPO 2009b

, E vans et al. 2008, H érar d et al. 2006 A noplophor a glabripennis (M otschulsky , 1853) A phytophagous China S outh-Central 2001, A T A T , DE, FR, IT FB, F A, I A cer , A esculus, B etula, _Carpinus, Fagus, P opulus, Salix Car ter et al. 2009,

Cocquempot 2007, Cocquempot et al. 2003, Dauber and M

itter 2001, EPPO 2004, H érar d et al. 2006, 2009 C allidiellum ru fi penne (M otschulsky , 1860) A phytophagous Eastern Asia, Japan

1906, FR ES, FR, IT FA, FB, G1, G5, J4 C upr essaceae (C upr essus macr ocarpa ) B ahillo and I turr ondobeitia 1995, Campadelli and S ama 1988, Cocquempot 2007 Chlor ophor us annularis (F abricius, 1787) A phytophagous Asia- Temperate 1991, ES ES G B amboo V iv es 1995 C yr thognathus forfi catus (F abricius, 1792) A phytophagous Africa 1872, MT MT U U nkno wn B er tolini 1872 D er olus mauritanicus B uquet, 1840 A phytophagous N or thern Africa 1884, FR ES ?, FR ? E7, F5, F8, FB, I2, X11 N erium oleander B rustel et al. 2002, F auv el 1884, M endizábal 1944, V er dugo 2004 T a b le 8.1.1.

List and characteristics of the Ceramb

ycidae species alien to E

ur

ope. S

tatus: A: Alien to E

ur

ope; C: cr

yptogenic species. Coun

tr y codes abbr eviations refer to ISO 3166 (s ee appendix I). H abitat abbr eviations r

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Longhorn beetles (Coleoptera, Cerambycidae). Chapter 8.1 215 F amily _Species Status R egime N ativ e range 1st r ecor d in E u rope In vaded countries H abitat H osts R efer ences D er oplia albida (B rullé, 1838) A phytophagous Canar y Islands 1988, ES ES E7, F6, FB, G5 Pelargonium V iv es 1995 L ucasianus lev aillantii (L ucas, 1846) A phytophagous N or thern Africa 1905, FR ES, FR, PT FA, G, FB C upr essus B rustel et al. 2002, Cocquempot et al. 2007, May et 1905, P ellegrin and Cocquempot 2001, P laza Lama 1990, V iv es 1995 N eoclytus acuminatus (F abricius, 1775) A phytophagous South- Central U.S.A.

1908, IT

CH, CZ, DE, FR, HR, HU, IT, ME, PT

-MAD, RS, SI FB, G, G1, G5, I2, X11 U lmus, Fr axinus, Juglans B ijaoui 1980, B rustel et al. 2002, Cocquempot 2007, H eyr ovský 1951, I lić 2005, P icar d 1937, P il and Stojano vić 2005, R eineck 1919, S ama 1984, T assi 1969, V illiers 1979, W inkler 1932, W ittenberg 2005 O xymer us aculeatus lebasi D upont, 1838 C phytophagous U nkno wn U nkno wn ES-CAN U C alophyllum Alluaud 1935 Pa randr a br unnea (F abricius, 1789) A phytophagous N or th America 1916, DE DE G, J1 T ilia, P opulus , deciduous trees G rämer 1961, N üssler 1961 P hor acantha r ecur va N ewman, 1840 A phytophagous A ustralia 1992, IT ES, GR, IL, IT , IT -SAR, IT -SIC, MT , PT G1 E ucalyptus B er cedo and B ahillo 1998, B er cedo and B ahillo 1999,

Černý 2002, Cocquempot 2007, Cocquempot and S

ama 2004, F riedman et al. 2008, M azz eo and S iscar o 2007, M ifsud 2002, M iquel 2008, O rousset 2000, P almeri and Campolo 2006, P ér ez M or eno 2001, R uiz and B arranco 1998,

Sama and Bocchini 2003, Sama et al. 2010,

W

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F amily _Species Status R egime N ativ e range 1st r ecor d in E u rope In vaded countries H abitat H osts R efer ences P hor acantha semipunctata (F abricius, 1775) A phytophagous A ustralia 1948, IL CY , FR,

FR-COR, ES, ES-CAN, GR, IL, IT

, IT -SAR, IT -SIC, MT , PT , PT -MAD FB, G, G1, G5, I2, X11 E ucalyptus B erger 1992, B rustel et

al. 2002, Cadahia 1980, Cav

alcaselle 1983, Černý

2002, Cocquempot 1993, Cocquempot 2007, Cocquempot and S

ama 2004,

M

ifsud and Booth 1997,

O rousset 1984, O rousset 1991, S ama et al. 2010, T eunissen 2002, V iv es 1995, W ang 1995 P hr yneta lepr osa (F abricius, 1775) A phytophagous South Tropical Africa

1997, FR FR, MT G M or us nigr a M ifsud and D andria 2002, V incent 2007 Taeniotes cay ennensis Th omson, 1859 A phytophagous Central America 1858, PT PT -AZ O U T ropical tr ees Sama 2006a T rinophylum cribr atum (B ates, 1878) A phytophagous India U nkno wn GB I2 D eciduous tr ees, Larix , P inus _{(polyphagous)} Du ff y 1953b , G ilmour 1948 X ylotr echus stebbingi G ahan, 1906 A phytophagous Central Asia 1990, IT CH, CY ,

DE, FR, GR, GR-CRE, GR-NEG, GR- SEG, IL, IT

, IT -SAR FB, G, G1, G5, I2, X11 A lnus, F icus, M or us, P opulus

Cocquempot 2007, Cocquempot and D

ebr euil 2006, D ioli and V igano 1990, K öhler 2000, S ama 2006b , Sama et al. 2010, Š efr ov á and Laštůvka 2005, T omicz ek and H oy er-T omicz ek 2008, W ittenberg 2005

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Longhorn beetles (Coleoptera, Cerambycidae). Chapter 8.1 217 F amily _species R egime N ativ e range In vaded countries H abitat H osts R efer ences A rhopalus rusticus (Linnaeus, 1758) phytophagous Continental Eur ope PT -AZ O, PT -MAD G3 P inus, P icea, A bies, Larix Fauv el 1897, P icar d 1937, S errano 1982 A romia moschata (Linné, 1758) phytophagous Continental Eur ope PT -AZ O I2 Salix, P opulus, A lnus Borges et al. 2005 C er amb yx carinatus K üster , 1846 phytophagous B alkans MT G P runus

Sama and Cocquempot 1986

C er amb yx nodulosus G ermar , 1817 phytophagous B alkans MT G P yr us, M alus Fauv el 1897, Schembri and S ama 1986 Clytus arietis (Linné, 1758) phytophagous Continental Eur ope PT -MAD E5, G, _{G1, G5} D eciduous tr ees (polyphagous) P icar d 1937, W ollaston 1854 G racilia minuta (F abricius, 1781) phytophagous Southern Eur ope A T , CH, , ES-CAN, IE, L V , LT , PT -AZ O, PT -MAD F3, G, G5 D eciduous tr ees (polyphagous Borges et al. 2005, B ytinski-S alz 1956, L ucht 1987, S peight 1988, W ollaston 1863 Icosium tomentosum atticum G anglbauer , 1881 phytophagous Southeastern Eur ope FR G3 C upr essaceae Cocquempot et al. 2007, P ellegrin 1990 M onochamus gallopr ovincialis (O livier , 1795) phytophagous Southw estern E ur ope NL G3 P inus D e F luiter 1950 M onochamus sar tor (F abricius, 1787) phytophagous N or thern E ur ope, Alps BE, , NL G3 P icea Fauv el 1884, W iel 1956, L ucht 1987 M onochamus sutor (Linnaeus, 1758) phytophagous Central and Nor thern E ur ope BE, PT G3 P icea, P inus Speight 1988, W ey ers 1876 M orimus asper funer eus M ulsant, 1863 phytophagous Southeastern Eur ope CZ, MT G D eciduous tr ees (polyphagous Schembri and S ama 1986, Š efr ov á and Laštůvka 2005 T a b le 8.1.2.

List and characteristics of the Ceramb

ycidae species alien

in E ur ope. Countr y codes abbr eviations r

efer to ISO 3166 (see appendix I). H

abitat abbr

e-viations r

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F amily _species R egime N ativ e range In vaded countries H abitat H osts R efer ences N athrius br evipennis (M ulsant, 1839) phytophagous Southw estern E ur ope A T , BE, CH, CZ, DE, GB, IE, L U, L V , PL, PT -AZ O F3 D

eciduous and conifer

tr

ees (polyphagous)

A

dlbauer 2006, Borges et al. 2005,

Du ff y 1953a, H eyr ovský 1930, K or cynski 1985, L ucht 1987, S liwinski 1958, S peight 1988, W eidner 1973, W ey ers 1875 P hymatodes testaceus (Linné, 1758) phytophagous Continental Eur ope PT -AZ O G D eciduous and fr uit tr ees, pr eferably on Q uer cus Fauv el 1897, P icar d 1937, W ollaston 1854 Poecilium lividum (R ossi, 1794) phytophagous Southeastern Eur ope BE, CH, CZ, DE, L U, NL G,J1 Q uer cus, C astanea L ucht 1987, H eyr ovský and S láma 1992, H orion 1974, Š efr ov á and Laštůvka 2005, W ittenberg 2005 Rhagium inquisitor (Linné, 1758) phytophagous Continental Eur ope IE G3 Conifers (P inus, P icea, A bies, Larix); deciduous tr ees ( B etula, F agus, Q uer cus ) Speight 1988 R osalia alpina (Linné, 1758) phytophagous Central Eur ope, Alps MT G, I2, J1 Fagus , and other deciduous tr ees H

orion 1974, Schembri and S

ama 1986 Stictoleptur a rubr a (Linné, 1758) phytophagous Central Eur ope PT -AZ O G 3 Conifers ( P inus, P icea, A bies, Larix ) Borges et al. 2005 St romatium unicolor (O livier , 1795) phytophagous Southeastern Eur ope PT -MAD G D eciduous tr ees

(mostly) and conifers (polyphagous)

Fauv el 1897, P icar d 1937 T richofer us fasciculatus (F aldermann, 1837) phytophagous Southeastern Eur ope CH, PT -MAD G D eciduous tr ees (polyphagous) Allenspach 1973, P icar d 1937 T richofer us griseus (F abricius, 1792) phytophagous Southeastern Eur ope CZ G F icus, P istacia, R osa Šefr ov á and Laštůvka 2005 X ylotr echus ar vicola (O livier , 1795) phytophagous Southeastern Eur ope SP-CAN G D eciduous tr ees (polyphagous) D emelt 1974